Memoirs of Museum Victoria 68:1-28 (2011) 
ISSN 1447-2546 (Print) 1447-2554 (On-line) 
http:// museum.com.au/About/Books-and-Journals/Journals/Memoirs-of-Museum-'Victoria 
A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae) 
Thomas A. Darragh 
Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia (tdarragh@museum.vic.gov.au) 
Abstract Darragh, Thomas A. 2011. A revision of the Australian fossil species of Zoila (Gastropoda: Cypraeidae). Memoirs of 
Museum Victoria 68: 1-28. 
Zoila Jousseaume, 1884 is a cowry genus, the species of which are now confined to the coastal waters of Western 
Australia and southern Australia. Six fossil species are known from southeast Australia, of which three are new, ranging 
in age from late Oligocene to middle Miocene, known from the Murray, Otway and Bass basins. Three fossil species are 
known from Western Australia, of which three are new, ranging in age from late Eocene to late Pliocene, known from the 
Carnarvon, Bremer and Eucla basins. Umbilia (Gigantocypraea) Schilder 1927 (type species Cypraea gigas McCoy) is 
regarded as a synonym of Zoila. The earliest record of the genus is Z. chathamensis (Cemohorsky, 1971), Paleocene/early 
Eocene, Chatham Islands, probably ancestral to the Australian fossils. Zoila is closely related to Cypraeorbis Conrad, 
1865, of which Bernaya Jousseaume, 1884 is regarded as a synonym. Living species are known to have no planktotrophic 
larval stage, so there is considerable variation in species morphology. Such direct development arose in these cowries at 
least as early as the late Eocene. Fossils described here are Zoila viathomsoni sp. nov., Z. didymorhyncha sp. nov., Z. 
glomerabilis sp. nov., Z. dolichorhyncha sp. nov., Z. mulderi (Tate, 1892), Z. platypyga (McCoy, 1876) (= consobrina 
McCoy, toxorhyncha Tate, platypyga simplicior Schilder), Z. gigas (McCoy, 1867) (= dorsata, gabrieli ), Z. campestris n. 
sp. and Z.fodinata n. sp. 
Keywords Gastropoda, Cypraeidae, cowries, Australia, Tertiary, new species, taxonomy 
Introduction 
Zoila is a member of a group of Australian cowries that 
includes Notocypraea, Umbilia and Austrocypraea, the 
members of which all seem to have direct larval development 
rather than having a planktotrophic larval stage as in most 
species of cowry (Wilson, 1985; Wilson and Clarkson, 2004). 
This paper reports research results that continue from a study 
of the Australian endemic genus Umbilia (Darragh, 2002). 
Subfamily classification 
The genus Zoila is usually placed in the subfamily Bernayinae 
Schilder, 1927 erected for the fossil genus Bernaya Jousseaume, 
1884, of which Zoilinae Iredale, 1935 is a synonym (Wilson 
and Clarkson, 2004). Meyer (2003), in his work on molecular 
systematics of cowries, retained most of the subfamilies 
previously used by many authors on the family, but did not 
place the genus Zoila in any subfamily (although his Table 3 
could imply a placement in the subfamily Bernayinae). 
However, following further molecular work, he subsequently 
(Meyer, 2004) placed the genera Zoila and Barycypraea 
within the subfamily Bernayinae. Cypraeorbinae Schilder, 
1927 was erected for the fossil genus Cypraeorbis Conrad, 
1865.1 consider that Bernaya is a synonym of Cypraeorbis, as 
argued below. Therefore, if subfamilies are regarded as useful 
in classification, Cypraeorbinae should be used instead. 
Bouchet and Rocroi (2005) included the subfamilies 
Bernayinae and Cypraeorbinae, along with three other 
subfamilies, within the synonymy of Gisortinae Schilder, 
1927, erected for the genus Gisortia Jousseaume, 1884. Species 
of Gisortia have an obsolete fossula, quite different from those 
of species of Cypraeorbis and Zoila, so this synonymy is 
unlikely. 
Nomenclatural history of Zoila fossil species 
The genus Zoila was established by Jousseaume (1884a) for 
three species of large living cowries from Western Australia. 
The first fossil species later assigned to this genus were 
described by Frederick McCoy from the Tertiary of Victoria. 
Cypraea gigas was described by McCoy in 1867 without 
illustration and in 1875, it was redescribed, figured and placed 
in the subgenus Aricia. Two more species, Cypraea (Aricia) 
platypyga and C. (A.) consobrina were described and figured 
by McCoy in 1876 and 1877, respectively. In 1890, Tate revised 
these three species and added another two species, C. 
toxorhyncha, regarded here as a synonym of Z. platypyga, and 
Cypraea dorsata, regarded here as a synonym of Z. gigas. In 
1892, Tate figured a new species C. mulderi and described it in 
1893, pointing out its affinities with C. platypyga. Tate 
assigned all these species to Cypraea. Harris (1897) assigned 
